The Question of a Presupracleithrum in Brachiopterygian Fishes

The presupracleithrum is an exoskeletal pectoral bone that occurs in Paleozoic and Mesozoic actinopterygian fishes. It has been equated more than once with an opercular element in brachiopterygian fishes. In recent cladistic analyses, this alleged homology is used to assign brachiopterygians to actinopterygians. However, a comparison of brachiopterygian and actinopterygian crania shows clearly that the former lack a presupracleithrum.     

Assessing metabolic constraints on the maximum body size of actinopterygians: locomotion energetics of Leedsichthys problematicus (Actinopterygii,Pachycormiformes)

Maximum sizes attained by living actinopterygians are much smaller than those reached by chondrichthyans. Several factors, including the high metabolic requirements of bony fishes, have been proposed as possible body‐size constraints but no empirical approaches exist. Remarkably, fossil evidence has rarely been considered despite some extinct actinopterygians reaching sizes comparable to those of the largest living sharks. Here, we have assessed the locomotion energetics of Leedsichthys problematicus, an extinct gigantic suspension‐feeder and the largest actinopterygian ever known, shedding light on the metabolic limits of body size in actinopterygians and the possible underlying factors that drove the gigantism in pachycormiforms. Phylogenetic generalized least squares analyses and power performance curves established in living fishes were used to infer the metabolic budget and locomotion cost of L. problematicus in a wide range of scenarios. Our approach predicts that specimens weighing up to 44.9 tonnes would have been energetically viable and suggests that similar body sizes could also be possible among living taxa, discarding metabolic factors as likely body size constraints in actinopterygians. Other aspects, such as the high degree of endoskeletal ossification, oviparity, indirect development or the establishment of other large suspension‐feeders, could have hindered the evolution of gigantism among post‐Mesozoic ray‐finned fish groups. From this perspective, the evolution of anatomical innovations that allowed the transition towards a suspension‐feeding lifestyle in medium‐sized pachycormiforms and the emergence of ecological opportunity during the Mesozoic are proposed as the most likely factors for promoting the acquisition of gigantism in this successful lineage of actinopterygians.     

Apertures of Craniate offactory Organs

The olfactory organs of craniates appear to derive from an anterior pair of erstwhile branchiothyria and their adjacent epidermal, neurogenic placodes. The organ is termed saccus nasalis if its source is the ectobranchial invagination of the branchiothyrium, and the saccus rhinalis if its source is the entobranchial evagination of the branchiothyrium. Nasal sacs occur in gnathostomes and lampreys, and rhinal sacs in hagfish. It is difficult, therefore, to avoid the conclusion that the olfactory organ has evolved more than once. The rhinal sac has only one aperture, the postica, which represents an internal branchiothyric orifice. Partial fusion of the right and the left rhinal sac led to the development of a postica communis. The nasal sac has either one, two or three apertures. Altogether there are nine different aperturae sacci nasalis, viz. the naris, the foris, the tremiscus, the two nariculae, the rimilla, the portula, the opiscus and the janua. The janua represents an external branchiothyric orifice and occurs in lampreys. The naris and the foris arose by subdivision of the orifice from which the janua emanated; they are found in osteolepipods. The naris is also found in urodeles, porolepiforms and some teleosts. The incurrent and excurrent naricules resulted from bipartition of a naris and are present in actinopterygians, coelacanthiforms, brachiopterygians, dipnoans and elasmobranchiomorphs. Of the four remaining openings of the nasal sacs the rimilla results from an inpushing of the epidermis, the portula from an outpocketing of the nasal sac, and the tremiscus from a combination of two such movements, the one inward and the other outward. Rimillae occur in cyprinodonts, anurans, urodeles, caecilians and porolepiforms, portulae in uranoscopids and bathydraconids, and tremisci in urodeles and porolepiforms. The opiscus, finally, is the opening through which the vomeronasal organ communicates with the oral cavity. It occurs in most lizards and snakes and probably also in some mammals.     

Olfactory organ of acipenseriformes and comparison with other actinopterygians: Patterns of diversity

The position and structure of the olfactory organ and its openings vary among actinopterygians. The anterior nasal opening is a simple perforation in the skin in many extant actinopterygians (e.g., acipenseriforms, lepisosteids, and primitive Recent teleosts) and represents the primitive condition. Polypterids and Amia each exhibit a derived condition, in which the anterior nasal opening extends into a tube. The olfactory organ is relatively far away from the anterior end of the elongate rostrum in acipenseriforms, whereas the olfactory organs are closer to the anterior end of the snout in extant actinopterygians (e.g., polypterids, lepisosteids, and amiids). In adults, olfactory organs are cuplike structures in most actinopterygians, but these organs are tubelike in polypterids. Among extant actinopterygians, a nasal diverticulum is present only in polypterids. Teleosts have accessory nasal sacs, but chondrosteans, polypterids, lepisosteids, and amiids lack them. The olfactory rosette is formed by primary folds or lamellae that may be placed anterior, lateral, posterior, and/or medial to the axis of the organ. Large acipenserids have 20–32 lamellae, polyodontids have 13–18 lamellae, lepisosteids have 8–10 lamellae, and Amia may have over 100. In teleosts, the number of lamellae varies from none or a few to over 200. Secondary lamellae are present in acipenseriforms, lepisosteids, and some advanced teleosts; secondary lamellae are interpreted as independently acquired in these lineages. Secondary lamellae are absent in Amia and primitive teleosts such as Elops and Hiodon. Tertiary lamellae are present in Acipenser oxyrhynchus. The arrangement of the primary lamellae in relation to the axis of the organ results in at least 11 patterns of the olfactory rosette in actinopterygians. Lamellae that are enclosed in a tubelike sac and that have an anteromedial diverticulum are specializations of polypterids. Primary lamellae anterior, lateral, and posterior to an elongate axis are characteristic of lepisosteids. The presence of primary lamellae lateral, medial, and posterior to an elongate olfactory axis is a synapomorphy of Halecomorpha (Amia plus teleosts). The absence of secondary lamellae is a synapomorphy of Halecomorpha. © 1994 Wiley-Liss, Inc.     

Morphology and Interrelationships of Primitive Actinopterygian Fishes

SYNOPSIS. The concept of the Actinopterygii as a natural groupof fishes was not generally accepted until early in this century.Ever since, the characterization of the group has been blurredby the problem of cladistian (polypterid) relationships. Froma review of the structure of polypterids and actinopts, it isconcluded that Cladistia are the sistergroup of Recent actinopterygians(Actinopteri), the two groups together comprising the Actinopterygii.Recent chondrosteans are more closely related to higher actinopts(Neopterygii) than to cladistians. The extinct Palaeonisciformesappear to be a paraphyletic group, comprising stem-group actinopterygians(e.g., Cheirolepts), stem-group actinopterans (e.g., Moythomasia)and relatives of higher actinopterans (e.g., Pteroniscus)     

Endoskeletal structure in Cheirolepis (Osteichthyes,Actinopterygii), An early ray‐finned fish

As the sister lineage of all other actinopterygians, the Middle to Late Devonian (Eifelian–Frasnian) Cheirolepis occupies a pivotal position in vertebrate phylogeny. Although the dermal skeleton of this taxon has been exhaustively described, very little of its endoskeleton is known, leaving questions of neurocranial and fin evolution in early ray‐finned fishes unresolved. The model for early actinopterygian anatomy has instead been based largely on the Late Devonian (Frasnian) Mimipiscis, preserved in stunning detail from the Gogo Formation of Australia. Here, we present re‐examinations of existing museum specimens through the use of high‐resolution laboratory‐ and synchrotron‐based computed tomography scanning, revealing new details of the neuro‐cranium, hyomandibula and pectoral fin endoskeleton for the Eifelian Cheirolepis trailli. These new data highlight traits considered uncharacteristic of early actinopterygians, including an uninvested dorsal aorta and imperforate propterygium, and corroborate the early divergence of Cheirolepis within actinopterygian phylogeny. These traits represent conspicuous differences between the endoskeletal structure of Cheirolepis and Mimipiscis. Additionally, we describe new aspects of the parasphenoid, vomer and scales, most notably that the scales display peg‐and‐socket articulation and a distinct neck. Collectively, these new data help clarify primitive conditions within ray‐finned fishes, which in turn have important implications for understanding features likely present in the last common ancestor of living osteichthyans.     

Observations of terrestrial locomotion in wild Polypterus senegalus from Lake Albert,Uganda

Polypterids, the most basal actinopterygians, are a group of fish long-considered living fossils and holding a key position for understanding fish and tetrapod evolution. Knowledge of the natural history of Polypterus is limited, their having been studied in little detail since the early 1900s. The locomotory habits of wild Polypterus senegalus from Lake Albert, Uganda, were investigated in 2014. High-speed videography demonstrated the capability of large Polypterus to move overland successfully. Contrary to previous evidence, field observations found that terrestrial locomotion in Polypterus is not inherently restricted by body size. Evidence that Polypterus exhibit this behaviour as part of their natural life history can be found in the existence of environmental challenges and the presence of adaptations for amphibious life.     

Early Pennsylvanian (Langsettian) fish assemblages from the Joggins Formation,Canada, and their implications for palaeoecology and palaeogeography

A review of all available specimens of fossil fishes from the classic Pennsylvanian Joggins locality of Nova Scotia, Canada, reveals the existence of a diverse community of chondrichthyans (xenacanthids, ctenacanthids and the enigmatic Ageleodus), acanthodians (gyracanthids), sarcopterygians (rhizodontids, megalichthyids, rhizodopsids and dipnoans) and actinopterygians (haplolepids). Reassessment of supposed endemic species (Ctenoptychius cristatus, Sagenodus plicatus, Gyracanthus duplicatus) indicates they are invalid, and overall, the assemblage comprises cosmopolitan taxa that were widespread around the coasts of tropical Pangaea. Strontium isotope analysis of fish remains and a critical study of their facies context suggest that these fish communities occupied bodies of water with salinities across the marine–freshwater spectrum. This preponderance of euryhaline forms implies a community structure quite distinct from that of today and might represent a transitory phase prior to the establishment of fully freshwater fish communities. Interpretation of fish ecology provides further evidence that the Joggins Formation was deposited in a paralic setting, and the recognition of one previously undetected brackish incursion strengthens the link between sedimentary cycles at Joggins and Milankovitch‐induced glacio‐eustatic change. Furthermore, interregional correlation of these marine transgressions supports palynostratigraphical arguments for an early Langsettian age for the Joggins Formation. This places tighter constraints on the age of the earliest known crown amniote, Hylonomus lyelli, an important calibration point used in phylogenomic studies.     

Development of cranial muscles in the actinopterygian fish Senegal bichir,Polypterus senegalus Cuvier, 1829

Polypterus senegalus Cuvier, 1829 is one of the most basal living actinopterygian fish and a member of the Actinopterygii. We analyzed the spatial and temporal pattern of cranial muscle development of P. senegalus using whole‐mount immunostaining and serial sectioning. We described the detailed structure of the external gill muscles which divided into dorsal and ventral parts after yolk exhaustion. The pattern of the division is similar to that of urodeles. We suggest that, the external gill muscles of P. senegalus are involved in spreading and folding of the external gill stem and the branches. The fibers of the external gill muscles appear postero‐lateral to the auditory capsule. In addition, the facial nerve passes through the external gills. Therefore, the external gill muscles are probably derived from the m. constrictor hyoideus dorsalis. In contrast to previous studies, we described the mm. interhyoideus and hyohyoideus fibers as independent components in the yolk‐sac larvae. The m. hyohyoideus fibers appear lateral to the edge of the ventral portion of the external gill muscles, which are probably derived from the m. constrictor hyoideus dorsalis. These findings suggest that the m. hyohyoidues is derived from the m. constrictor hyoideus dorsalis in P. senegalus . In other actinopterygians, the m. hyohyoideus is derived from the m. constrictor hyoideus ventralis; therefore, the homology of the m. hyohyoidues of P. senegalus and other actinopterygians remains unclear. J. Morphol. 278:450–463, 2017. © 2017 Wiley Periodicals, Inc.     

CT scan reconstruction of the palate region of Latimeria chalumnae

Synopsis The palate of Latimeria chalumnae is described based mainly on three-dimensional CT scan reconstruction. It is compared with that of other osteichthyans. The palate of L. chalumnae compares best with that of rhipidistians; it is more advanced than that of actinopterygians in having fewer bones. This tendency toward bone reduction in the palate is even more pronounced in dipnoans. The interpretation of features of the Early Devonian genus Diabolepis determines if authors consider dipnoans or actinistians more closely related to tetrapods. Both groups are only distant relatives of tetrapods.     

Reconstruction of cranial and hyobranchial muscles in the triassic temnospondyl Gerrothorax provides evidence for akinetic suction feeding

The cranial and hyobranchial muscles of the Triassic temnospondyl Gerrothorax have been reconstructed based on direct evidence (spatial limitations, ossified muscle insertion sites on skull, mandible, and hyobranchium) and on phylogenetic reasoning (with extant basal actinopterygians and caudates as bracketing taxa). The skeletal and soft‐anatomical data allow the reconstruction of the feeding strike of this bottom‐dwelling, aquatic temnospondyl. The orientation of the muscle scars on the postglenoid area of the mandible indicates that the depressor mandibulae was indeed used for lowering the mandible and not to raise the skull as supposed previously and implies that the skull including the mandible must have been lifted off the ground during prey capture. It can thus be assumed that Gerrothorax raised the head toward the prey with the jaws still closed. Analogous to the bracketing taxa, subsequent mouth opening was caused by action of the strong epaxial muscles (further elevation of the head) and the depressor mandibulae and rectus cervicis (lowering of the mandible). During mouth opening, the action of the rectus cervicis muscle also rotated the hyobranchial apparatus ventrally and caudally, thus expanding the buccal cavity and causing the inflow of water with the prey through the mouth opening. The strongly developed depressor mandibulae and rectus cervicis, and the well ossified, large quadrate‐articular joint suggest that this action occurred rapidly and that powerful suction was generated. Also, the jaw adductors were well developed and enabled a rapid mouth closure. In contrast to extant caudate larvae and most extant actinopterygians (teleosts), no cranial kinesis was possible in the Gerrothorax skull, and therefore suction feeding was not as elaborate as in these extant forms. This reconstruction may guide future studies of feeding in extinct aquatic tetrapods with ossified hyobranchial apparatus. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

The Embryonic and Larval Development of Polypterus senegalusCuvier, 1829: its Staging with Reference to External and Skeletal Features, Behaviour and Locomotory Habits

The embryonic and larval development of the Polypteriformes, the presumed sister group of all other living actinopterygians, is poorly known. The main reason is the scarcity of successful breedings in captivity and therefore the lack of developmental series of any one polypterid species. A series of five successful breedings of P. senegalusnow makes it possible to define developmental stages of this species based on numerous closely timed specimens. The staging given here focuses on external embryonic and larval features: epidermal surface structures documented by SEM, colour pattern, development of fins and squamation, larval feeding and locomotory behaviour. The development of P. senegalusis characterized by a long free embryonic phase. Suction feeding is performed from the beginning of larval life (apterolarval phase). The pectoral fins start to become employed for slow locomotion and as supportive structures at around the same time. Olfactorily guided prey capture, however, is observed later in the pterolarval phase. Quantitative kinematic data also demonstrate a change in the mode of undulatory locomotion during this phase. Sustained axial undulation becomes confined to the posterior abdominal and caudal region of the body. At about the same time the paraxial high frequency undulation of the pectoral fin fold is replaced by the characteristic propeller-like movement of much greater amplitude and wavelength. Surfacing for aerial breathing is not seen before a marked change in colouration has taken place at the beginning of the juvenile period. The external gills slowly become reduced during this period. The definitions of larval and juvenile stages given here may advance understanding of developmental processes in the ontogeny of these primitive actinopterygians, and may serve as a tool for comparison with the ontogeny of Tetrapoda and Dipnoi, as well as to that of some “primitive” groups of Actinopterygii. Judging from its distribution among extant taxa, embryonic and larval ciliation is a character that most probably belongs to the grundplan? of Osteognathostomata. Phylogenetic evaluation is not so clear for the two other prominent embryonic and larval specializations found in Polypterus: upper labial attachment glands and opercular external gills. © 1997 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd.     

Homology of the Fifth Epibranchial and Accessory Elements of the Ceratobranchials among Gnathostomes: Insights from the Development of Ostariophysans

Epibranchials are among the main dorsal elements of the gill basket in jawed vertebrates (Gnathostomata). Among extant fishes, chondrichthyans most resemble the putative ancestral condition as all branchial arches possess every serially homologous piece. In osteichthyans, a primitive rod-like epibranchial 5, articulated to ceratobranchial 5, is absent. Instead, epibranchial 5 of many actinopterygians is here identified as an accessory element attached to ceratobranchial 4. Differences in shape and attachment of epibranchial 5 in chondrichthyans and actinopterygians raised suspicions about their homology, prompting us to conduct a detailed study of the morphology and development of the branchial basket of three ostariophysans (Prochilodus argenteus, Characiformes; Lophiosilurus alexandri and Pseudoplatystoma corruscans, Siluriformes). Results were interpreted within a phylogenetic context of major gnathostome lineages. Developmental series strongly suggest that the so-called epibranchial 5 of actinopterygians does not belong to the epal series because it shares the same chondroblastic layer with ceratobranchial 4 and its ontogenetic emergence is considerably late. This neomorphic structure is called accessory element of ceratobranchial 4. Its distribution among gnathostomes indicates it is a teleost synapomorphy, occurring homoplastically in Polypteriformes, whereas the loss of the true epibranchial 5 is an osteichthyan synapomorphy. The origin of the accessory element of ceratobranchial 4 appears to have occurred twice in osteichthyans, but it may have a single origin; in this case, the accessory element of ceratobranchial 4 would represent a remnant of a series of elements distally attached to ceratobranchials 1–4, a condition totally or partially retained in basal actinopterygians. Situations wherein a structure is lost while a similar neomorphic element is present may lead to erroneous homology assessments; these can be avoided by detailed morphological and ontogenetic investigations interpreted in the light of well-supported phylogenetic hypotheses.     

The anatomical components of the cardiac outflow tract of chondrichthyans and actinopterygians

The outflow tract of the fish heart is the segment interposed between the ventricle and the ventral aorta. It holds the valves that prevent blood backflow from the gill vasculature to the ventricle. The anatomical composition, histological structure and evolutionary changes in the fish cardiac outflow tract have been under discussion for nearly two centuries and are still subject to debate. This paper offers a brief historical review of the main conceptions about the cardiac outflow tract components of chondrichthyans (cartilaginous fish) and actinopterygians (ray‐finned fish) which have been put forward since the beginning of the nineteenth century up to the current day. We focus on the evolutionary origin of the outflow tract components and the changes to which they have been subject in the major extant groups of chondrichthyans and actinopterygians. In addition, an attempt is made to infer the primitive anatomical design of the heart of the gnathostomes (jawed vertebrates). Finally, several areas of further investigation are suggested. Recent work on fish heart morphology has shown that the cardiac outflow tract of chondrichthyans does not consist exclusively of the myocardial conus arteriosus as classically thought. A conus arteriosus and a bulbus arteriosus, devoid of myocardium and mainly composed of elastin and smooth muscle, are usually present in cartilaginous and ray‐finned fish. This is consistent with the suggestion that both components coexisted from the onset of the gnathostome radiation. There is evidence that the conus arteriosus appeared in the agnathans. By contrast, the evolutionary origin of the bulbus is still unclear. It is almost certain that in all fish, both the conus and bulbus develop from the embryonic second heart field. We suggest herein that the primitive anatomical heart of the jawed vertebrates consisted of a sinus venosus containing the pacemaker tissue, an atrium possessing trabeculated myocardium, an atrioventricular region with compact myocardium which supported the atrioventricular valves, a ventricle composed of mixed myocardium, and an outflow tract consisting of a conus arteriosus, with compact myocardium in its wall and valves at its luminal side, and a non‐myocardial bulbus arteriosus that connected the conus with the ventral aorta. Chondrichthyans have retained this basic anatomical design of the heart. In actinopterygians, the heart has been subject to notable changes during evolution. Among them, the following two should be highlighted: (i) a decrease in size of the conus in combination with a remarkable development of the bulbus, especially in teleosts; and (ii) loss of the myocardial compact layer of the ventricle in many teleost species.     

An analysis of nucleotide and amino acid variability in the barcode region of cytochrome c oxidase I (cox1) in fishes

The 655 bp cytochrome c oxidase subunit I barcode region of single specimens of 388 species of fishes (four Holocephali, 61 Elasmobranchii and 323 Actinopterygii) was examined. All but two (Urolophus cruciatus and Urolophus sufflavus) showed different cox1 nucleotide sequences (99.5% species discrimination); the two that could not be resolved are suspected to hybridize. Most of the power of cox1 nucleotide sequence analysis for species identification comes from the degenerate nature of the genetic code and the highly variable nature of the third codon position of amino acids. Variation at the third codon position is bimodally distributed, and the more variable mode is dominated by amino acids with four or six codons, while the less variable mode is dominated by amino acids with two codons. The ratio of nonsynonymous to synomymous changes is much less than one, indicating that this gene is subject to strong purifying selection. Consequently, cox1 amino acid sequence diversity is much less than nucleotide sequence diversity and has very poor species resolution power. Fourteen of the 16 amino acid residues recognized as having important functions in the region of cox1 sequenced were completely conserved over all 388 species (and the bovine cox1 sequence), with one fish species varying at one of these sites, and three fish at another site. No significant differences in amino acid conservation were observed between residues in helices, strands and turns. Patterns of nucleotide and amino acid variability were very similar between elasmobranchs and actinopterygians.     

Development of the muscles associated with the mandibular and hyoid arches in the Siberian sturgeon,Acipenser baerii (Acipenseriformes: Acipenseridae)

The skeleton of the jaws and neurocranium of sturgeons (Acipenseridae) are connected only through the hyoid arch. This arrangement allows considerable protrusion and retraction of the jaws and is highly specialized among ray‐finned fishes (Actinopterygii). To better understand the unique morphology and the evolution of the jaw apparatus in Acipenseridae, we investigated the development of the muscles of the mandibular and hyoid arches of the Siberian sturgeon, Acipenser baerii. We used a combination of antibody staining and formalin‐induced fluorescence of tissues imaged with confocal microscopy and subsequent three‐dimensional reconstruction. These data were analyzed to address the identity of previously controversial and newly discovered muscle portions. Our results indicate that the anlagen of the muscles in A. baerii develop similarly to those of other actinopterygians, although they differ by not differentiating into distinct muscles. This is exemplified by the subpartitioning of the m. adductor mandibulae as well as the massive m. protractor hyomandibulae, for which we found a previously undescribed portion in each. The importance of paedomorphosis for the evolution of Acipenseriformes has been discussed before and our results indicate that the muscles of the mandibular and the hyoid may be another example for heterochronic evolution.     

Paired fin skeletons and relationships of the fossil group Porolepiformes (Osteichthyes: Sardcopterygii)

The endoskeletal girdles, anocleithrum and paired fin supports of the porolepiform fish Glyptolepis (Osteichthyes: Sarcopterygii: Porolepiformes) are figured and described. The pectoral fin skeleton is known from the proximal part only and the pelvic fin skeleton is fragmentary, but the scapulocoracoid, anocleithrum and pelvic girdle can be reconstructed in their entirety. The anocleithrum is entirely subdermal. The pectoral fin skeleton in shown to be biserial, with a large number of axial mesomeres, whereas the pelvic fin contains fewer mesomeres and is strongly asymmetrical with very few postaxial radials. The scapulocoracoid is essentially similar to a reconstruction figured by Jarvik (1980), but has a more elongate glenoid; this has functional implications. The pelvic girdle consists of two separate halves as in Eusthenopteron, but differs from that genus in lacking dorsolateral rami. A brief survey of the evidence of paired fin structure in other porolepiform genera is carried out to establish whether the structures seen in Glyptolepis are likely to be representative for the Porolepiformes. A study of the morphology and muscle attachments of the paired fin skeletons indicates that the pattern of fin movement was significantly different from that in Neoceratodus. The fin supports and girdles of Glyptolepis are compared with those of other sarcopterygian groups as well as with actinopterygians, placoderms and sharks, in order to establish evolutionary polarities. Glyptolepis is shown to display a number of derived characters. The information gained from the comparison is used to construct a maximum parsimony cladogram, which places coelacanths as the sister group of porolepiforms + lungfishes, with the rhizodonts + tetrapods and osteolepiforms as successive sister groups of this clade. Characters of uncertain polarity are considered in the light of this cladogram. A comparison with recently published cladograms shows that none are completely compatible with the results from this study.